Fossil from the Bemhridge Marls, Palaeogene of the Isle of Wight, southern England

E. A. J arzembowski Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7 SBD Contents Synopsis 238 Introduction 239 Previous studies 240 Notes on lithology, inclusions and method of study 240 Field sections . 242 Correlation 243 Systematic descriptions 244 Order Isoptera 244 Family Mastotermitidae 244 Mastotermes Froggatt 244 Mastotermes anglicus von Rosen 245 Family Kalotermitidae 249 Genus Kalotermes Hagen 249 Kalotermes disruptus (Cockerell) 249 Family Rhinotermitidae 251 Genus Reticulitermes Holmgren 251 Reticulitermes sp. 251 Family Termitidae 253 A . 253 Family uncertain 253 Order Plecoptera . 253 Family Leuctridae 253 Genus Leuctra Stephens 253 Lellctra priscula (Cockerell) 253 Order Neuroptera 255 Family Mantispidae . 255 Genus Promantispa nov. 255 Promantispa relicta (Cockerell) 255 Family Hemerobiidae 257 Genus Hemerobius Linne 257 Hemerobius tinctus sp. nov. 257 Genus Nellronema McLachlan 259 ? NeurOflema sp. A 259 '! Nellronema sp. B 259 Genus indet. 260 Species A 260 Species B . 260 Species C . 260 Family Chrysopidae 260 Species A . 260 Family Sisyridae 261 Species A . 261 Family uncertain 262 Order Mecoptera . 262 Family Bittacidae 262

8ull. Br. lv/us. nat. Hist. (Geol.) 33 (4) : 237-293 Issued 27 March 1980 238 E. A. JARZEMBOWSKI Genus Bittacl/s Latreille . 262 Bittacils vetertillS (Cockerell) 262 Bitfacus sp. A 263 Order 263 Family 263 Genus Micropterix Hubner 263 ft,licropterix anglica sp. nov. 263 Family 265 Genus Prohepialus Piton 265 Prolrepialus sp. 265 Family 267 Genus Paratriaxomasia nov. 267 Paratriaxomasia solentensis sp. nov. 267 Genus indet. 269 Species A . 269 SuperfamilY 269 Species A . 269 Family 270 Genus Copromorpha Meyrick 270 Copromorpha fossilis sp. nov. 270 Microlepidoptera, Family uncertain 270 Species A-L 270-275 ? Family 275 Genus Gurtletia Cockerell 275 Glirnetia durranti Cockerell 275 Family 276 Genus pyralites Heer 276 Pyralites preecei sp. nov. 276 Family Geometridae . 278 Genus Geometridites Clark et al. 278 Geometridites larentiiformis sp. nov. 278 Family 279 Genus Nymphalites Scudder 279 Nymphalites zellneri sp. nov. 279 Family 282 Genus Lithopsyehe Butler 282 Lithopsyehe ant/qua Butler 283 ? Superfamily 284 Genus indet. 284 Conclusions 284 Acknowledgements 286 References 286 Index 291

Synopsis The palaeontology, sedimentology and stratigraphy of the Bed (Bembridge Marls) are discussed and previous work summarized. Five insect orders are revised, notes on the biology of living relatives are included and the palaeoecology considered. lsoptera: Mastotermes angliclls von Rosen is redescribed, Sisyra (?) disrupta Cockerell is transferred from the Nellroptera to Kalotermes s. lat. (Kalotermitidae); a species of Reticulitermes (Rhinotermitidae) and a termitid are described but not named. The Isoptera suggest a warm subtropical-tropical climate. Plecoptera: Nemoura priscufa Cockerell is transferred to the genus Leuctra (Lellctridae). Megaloptera: the record of Raphidia is shown to be erroneous. Neurop­ tera: Mantispidae: Mantispa reliefa Cockerell is transferred to Promantispa gen. nov. Hemerobiidae: Hemerobius tinclus sp. nov., ? Neuronema spp. A and B, and 3 unnamed species are described. Chry­ sopidae: a species of Chrysopinae (s. lat.) is described but not named. The affinities of an extinct sisyrid are discussed. Mecoptera: Panorpa vetertla Cockerell (Panorpidae) is transferred to the genus Blttacus (Bittacidae); an unnamed species of BiftacliS is described. Lepidoptera: lvlicropterix anglica sp. nov. PALAEOGENE INSECTS 239 (Micropterigidae), Paratriaxomasia solentensis gen. et sp. nov. (Tineidae), Copromorpha fossilis sp. nov. (Copromorphidae), Pyralites preecei sp. nov. (Pyralidae), Nymphalites zeuneri sp. nov. (Nymphalidae), Prohepialus sp. (HepiaJidae) and Ceometridites larentiiformis sp. nov. (Geometridae) are described. Lithopsyche antiqua Butler is transferred to the Lycaenidae and the placement of Curt/etia durranti Cockerell in the Cossidae is questioned. A tineid, a tentative gelechioid, 12 other species of microJepid­ optera and a possible papilionoid are described but not named.

Introduction

The Bembridge Marls are a predominantly argillaceous formation, 21~36 m thick, preserved only in the northerly half of the Isle of Wight in southern England (Fig. 1). Near the base of the formation is a thin clay bed, generally less than a metre thick, with concretions and tabular bands of fine-grained argillaceous limestone and hard marl. The latter, on the NW coast near Cowes, have been noted for their insect remains for just over a century and constitute the 'Insect Lime­ stone' of authors (Daley 1973a). However, the calcareous and argillaceous developments together

1

Hamstead Ledge

/

Cliff End

~ __~_~~5 Km

Fig. 1 Outcrop map of the Bembridge Marls. form a distinct unit (ibid.) and the name Insect Bed (Bembridge Marls) used by Curry (1958) for a single limestone is here applied collectively. This horizon has yielded the only sizeable insect fauna in the British Tertiaries above the Lower Eocene, some fifteen orders being repre­ sented. The fauna also includes molluscs, an ostracod, an anostracous crustacean, an isopod, () and avian remains (feathers, Fig. 61, p. 268), the last three of which are scarce. Although named after the insects, the bed is probably better known in the palaeonto­ logical literature for its flora due to the work of Reid & Chandler (1926). However, apart from the leaves of a reed, well-preserved plant macrofossils are rare. Insect fragments are fairly frequent, and richer pockets are sometimes encountered (Jarzembowski 1976). The most extensive collections from this bed are now preserved in the British Museum (Natural History). These were largely collected by E. J. A'Court Smith (l814~1900), collecting between West Cowes and the Newtown River from about 1859 (Smith 1874). Apparently he discovered the insects following a comment by Forbes (1856 : 58~59) on the insect-bearing nature of some similar limestones in the Purbeck Beds (late Jurassic~early Cretaceous) on the mainland (Hughes 1922 : 67; Jackson 1933). Much of A'Court Smith's material subsequently went to R. W. Hooley and the Rev. P. B. Brodie, and the history of the collections and their acquisition by the Museum are given in Reid & Chandler (1926) and Crane & Getty (1975). Notable smaller collections from the Insect Bed include one at the Museum of Isle of Wight Geology, which was mainly the work of G. W. Colenutt, and another consisting of sonic of Brodie's material in the Lacoe Collection at the United States National Museum; I have myself collected from the bed since 1966. The fossil insect collections at the British Museum (Natural History) contain over 3900 pieces of 'Insect Limestone' of which a large proportion bear two or more specimens. However, a 240 E. A. JARZE,'vIBOWSKI number of these are indeterminate and some species, especially in the Formicidae, are repre­ sented by numerous examples. The present paper is confined to the Isoptera, Plecoptera, Neurop­ tera, Mecoptera and Lepidoptera, which although rare or uncommon have yielded a varied range of taxa; several other orders are being studied. AU available material is documented, and in addition to systematic and taxonomic considerations, the ecological and biological require­ ments of Recent relatives are briefly discussed.

Previous studies A'Court Smith (I874) appears to be the first published record of the occurrence of remains in the Insect Bed near Cowes. Woodward (1878, 1879) published a list of identifications made by Frederick Smith of the Department of Zoology, British Museum (Natural History), based on a sample of A'Court Smith's collection. In the 1879 paper Woodward described an anostracan, Branchipodites vectcnsis, and an isopod, Eosphaeroma fluviatile, from the same horizon; Martini (1972) showed the latter to be a synonym of the continental species Eosphaeroma margarum (Desmarest 1822). McCook (1888a, b) gave two identical descriptions of a mygalo­ morph , Eoatypus woodwardii, and Jones & Sherborn (1889) described an ostracod Pota­ mocypris brodiei which Haskins (1968) compared with Cypridopsis, another freshwater genus. The first description of an insect, Lithopsyche antiqua Butler 1889, was accompanied by c. colour plate. This was followed by von Rosen (1913) on fossil M~astotermes, and in 1915, T. D. A. Cockerell described 33 new insect species from the Lacoe Collection. He subsequently made selective studies of the main collection at the BM(NH): Cockerell (1917a, 1921a, b, c, 1922); Cockerell & Andrews (1916); Cockerell & Haines (1921). The Formicidae were studied by Donisthorpe (1920), the Culicidae by Edwards (1923) and Orthoptera by Zeuner (1939). These constitute the main works on the insect fauna and, although the flora was monographed by Reid & Chandler (1926), there is no comprehensive work on the . Chandler (1964) reviewed the Bembridge Flora, including the microfloral work of Machin (nee Pallot) published later in 1971. Although the junction of the Bembridge Marls and Bembridge Limestone is unconformable (Daley & Edwards 1971), the palaeobotanists considered the Bembridge Beds to contain a single flora, and it is not always clear which taxa occur in the Insect Bed. Daley (1969) studied the palaeoenvironment of the Bembridge Marls, and (1971, 1972a) was specifically concerned with sedimentary deformational structures in the 'Insect Limestone'. Jarzembowski (1976) first recorded insects in this bed on the east side of the island near St Helens, 18 km from CO\ves.

Notes on lithology, inclusions and method of study The insect-bearing lithology is usually a fine-grained argillaceous limestone with a conchoidal fracture and blue-grey colouration when fresh. It weathers externally producing a distinctive brownish yellow rind, and on broken surfaces the weathered zone has a sharp junction with the unweathered material. Older pebbles, especially near West Cowes, may only have a small region of 'blue' near the centre, and the weathered zone occasionally shows concentric colour banding. At outcrop it occurs in dark blue-grey clay as oval con;;retions, up to 20 cm thick, or as thinner tabular bands, and can be of a more marly constitution; thin, laterally continuous shell bands delimit the bed (Figs 2-4, p. 242). The calcareous developments tend to occur in courses within the clay more or less parallel to the bedding, which is of local stratigraphic value. There is a definite arching of the upper shell bed over large concretions, but in some places the undulations are less easy to explain. Silty laminae, 1-4 mm thick and 3-90 mm apart, are usually evident in the limestone, marked by finely-comminuted carbonaceous detritus. They appear to have no vertical regularity and erosional bases are occasionally evident. The laminae may contain ostracod valves, fragments of drifted vegetation -- the only common recognizable ones being of TypJza latissima Braun - and insect wings and body fragments; groups of shells belonging to the freshwater genus Galba some- PALAEOGENE INSECTS 241 times occur. The freshwater genera Hippeutis and Vivip.2rus also occur but are infrequent. Plant fossils are usually accompanied by a brown carbonaceous residue, and small black fragments of vegetable material are occasionally present. One of the latter has been examined and identified as natural fusain by Professor T. M. Harris. The limestone has yielded some small grains of resin: one of these (In. 17436) was analysed by infrared spectroscopy, but the spectra obtained were too weak for comparison with those of other Tertiary resins. The laminae occasionally yield concentrates of articulated insect bodies, often associated with abundant, complete examples of Branehipodites vee tensis , but such rich developments appear to be very local. Salt pseudomorphs, comminuted shell debris, seeds, spiders and portions of bird feathers are rare: the last are the only vertebrate fossils found. Insect bodies and wings are normally preserved in good relief, although substantial compression is evident in some of the more argillaceous laminae; well-preserved insects as well as fragments occur in the interbedded limestone, but there is never any concentration of material. Chitin is absent only from weakly sc1erotized areas and the colour pattern may be well preserved: comparison with related forms in the living fauna indicates that some colours are close to those in life (pp. 259, 262). The body cavities are often infilled with a pink calcareous material which obscures structures, but external moulds in the enclosing matrix show very fine detail (Fig. 59, p. 267) and the 'Insect Limestone' has been justifiably referred to as 'opaque amber'. The insects are predominantly imagines of small to medium size. The preservation of relief and soft-bodied, immature stages (Fig. 62, p. 268) as well as the fragile Branehipus-like crustacean, together with the arching of the upper shell bed, indicate early lithification of the matrix, but the occasional penetration of burrows from above, and development of the small-scale sedimentary deformation described by Daley (1971, 1972a) and evident in the limestone at all the main localities, suggest that it was not immediate. Pantin (1958) provides evidence for the length of time involved in the formation of diagenetic calcareous concretions. It is likely that the limestone was formed within several thousand years of deposition of the sediment, and perhaps less judged by the preservation of the soft-bodied arthropods. I have not observed any insect macrofossils in the clays adjacent to fossiliferous concretions although a similar lamination is evident on fresh surfaces. The remarks so far are based on the north-western localities. Whilst the lithological comments apply equally well on the east coast, there are some differences in the biota. St Helens and Priory Bay have yielded five orders of insects but only a few more complete specimens and the occasional ostracod and plant fragment. The author has only collected here since late 1975 and it is likely that more will be found. However, sustained search near Bembridge Foreland has yielded only a single insect specimen (see p. 243). Whilst the calcareous part of the Insect Bed is well developed and can be traced south into Whitecliff Bay, no fossils were found except for two fragments of lymnaean shell near the middle of Howgate Bay. The fossiliferous localities for insects are there­ fore confined to the northern part of the outcrop area of the Bembridge Marls (Fig. I). Insect remains do occur at other levels in the formation, but very few specimens have been found to date. The A'Court Smith collection includes examples in a clay-ironstone matrix from the north-west coast, and I have seen one or two fragments in the sideritic ironstone concretions from the upper part of the Bembridge Marls littering the shore near Whitecliff Bay Point (Daley 1973b). Where the limestone is exposed in the lower part of a cliff or slump and is accessible to marine erosion, it is usually well represented amongst the blocks and pebbles of the upper part of the shore, and splitting of this debris can be more rewarding. A hand-lens is necessary for examining fractured surfaces and small forms are more readily observed in oblique light. Specimens are rarely exposed properly owing to the curved fracture of the rock and both halves may show unique features. All the necessary preparation for this study was done by the author under a microscope with an electric engraver (vibrotool) or a tungsten needle. Routine examination of rock specimens under the microscope (x 16) also revealed a number of forms previously un­ noticed. The examination of chitinous material was often facilitated by immersing the specimen in liquid to heighten contrast; as the matrix may contain salt, especially the softer limestone and 242 E. A. JARZEMBOWSKI hard marl, aqueous solutions were avoided and xylene was used for short examinations and paraffin for longer periods. Dry examination was also aided by using oblique lighting from several directions and referring to modern specimens of related forms.

Field sections Natural exposures of the Insect Bed are confined to coastal sections and there appear to be no inland records from artificial excavations. The sections were examined between summer 1976 and spring 1977.

North-west coast. The 'Insect Limestone' is developed here only to the east of the Newtown River (Fig. 1, p. 239). The Insect Bed is represented by a thin band of clay just west of the river behind Hamstead Ledge and appears to be absent at Cliff End (Daley 1973a). Across the river, the strata descend gently on the eastern limb of the Porchfield anticline and the bed is seen at 433927\ a short distance before the western end of Saltmead Ledge. It is exposed in the lower part of the cliff for some 500 m as far east as 438928 and fossiliferous blocks occur on the shore. The bed is 0·65-0,85 m thick and four calcareous courses are recognizable (Fig. 2). The uppermost includes some large concretions and may account for the thinness of the course below. At 440930 two blocks of limestone were seen apparently in situ in the upper part of the foreshore. Pebbles of 'Insect Limestone' die out on the beach a short distance east of Salt mead Ledge (= Thorness Point/Ledge).

------_. - -- ._-- - -- ~ -- _._- - ~ ------'- --'-- _.. ------.------

o

- - ---'- - -,~ --. ------'- - -­ . -- - --.. ------'-"------==~~g~c:..:=~:?=""b BW III ===~===.o:n... O-SO-O-S5m BWI-II 0-17 m above Bembridge Limestone Figs 2-4 Sections of the basal part of the Bembridge Marls. Fig. 2, near Saltmead Ledge. Fig. 3, immediately south of Gurnard Ledge. Fig. 4, near St Helens. (Black: limestone. Oblique shading: hard marl. Dashed lines: laminated marly clay. Bivalve: Corbiculidae. Small gastropod: Tarebia. Large gastropod: Pofamides. The bed numbers to the right of Figs 2-3 are after Daley 1973a).

The bed is next seen on the east side of Thorness ( = Thorney) Bay, where a limestone course outcrops on the upper part of the foreshore behind Sticelett Ledge (461941). The bed gradually rises from the shore north towards Gurnard Ledge (= Gurnard Point) on the northern limb of the Thorness syncline and is about 2 m above cliff base behind the latter (463946). It is best exposed in the low cliff for some 140 m south of the ledge and is 0,65-0,75 m thick. Two calcareous courses are developed: a lower more or less continuous tabular band of limestone (Insect Bed of Curry 1958) and an upper discontinuous band of concretions, often less well cemented (Fig. 3). The clay yields sporadic examples of worn Po tam ides and corbiculid valves, rare examples of the latter also being found in the calcareous part. Between Gurnard Ledge and Gurnard (= Gurnet) Bay there is a kilometre of tumbled cliff line in the Bembridge Beds but there are no further

1 National Grid Reference, 100 km square 40 (SZ), as are the subsequent references. PALAEOGENE INSECTS 243 sections in the Insect Bed on the northern coast of the island before St Helens. Some character­ istic limestone was however found at several points amongst the slips 500-800 m NE of Gurnard Ledge, rising towards Gurnard from approximately 6 to 11 m above cliff base. Pieces of fossili­ ferous limestone occur amongst the shore debris as far as West Cowes (490966). A'Court Smith worked along the coast from West Cowes to the Newtown River (White 1921) although the specimens are generally labelled 'Gurnet (or Gurnard) Bay'. The most important collecting site was on the east side of Thorness Bay where the limestone outcropped on the shore (Brodie 1878, Reid 1889), which suggests that it was at Sticelett Ledge, and the latter has recently produced some relatively well-preserved material (Jarzembowski 1976). G. W. Colenutt refers to the site as Sticelett in correspondence (ibid.) but an accompanying sketch map by the same author is less helpful.

East Coast. Commencing in the north, a section of the Insect Bed is found NE of St Helen's Church (Fig. 4), 200-300 m south of Node's Point (638900). Although numerous blocks of limestone occur in the SW corner of the adjacent Priory Bay, no sections were observed amongst the slips. At the Point the bed is 9 m above the base of the cliff, and 0·7 m at the southern end of the section as the strata dip towards the axis of the Bembridge syncline. It is 0,60-0·85 m thick with a single course of limestone concretions. The lower shell bed is only well-developed in the southern part of the section. Continuing south, the Insect Bed is next seen near Bembridge Foreland, 2·7 km SE of St Helens, where a limestone course outcrops on the upper part of the foreshore amongst the beach cover and can be traced into Howgate Bay. At 654872 the limestone is 15 em thick and yielded a single Oecophylla wing. Further south the bed is seen in the lower part of the cliff near Whitecliff Bay Point, gradually ascending to the clifftop in Whitecliff Bay: the limestone becomes replaced by a more marly course, and there are no insect remains.

Correlation Edwards (1966) gives a summary of the stratigraphy of the younger Palaeogene strata of the Hampshire Basin and an extensive bibliography is found in Edwards (1971). The Bembridge Marls are near to the Eocene-Oligocene boundary, but different workers have drawn it variously above and below the formation. Edwards (1966) followed Curry (I958) who regarded the Bembridge Marls as of Oligocene, Lattorfian age, but as can be seen from Table 1, there is still no general agreement on the position of the boundary. The Insect Bed (Bembridge Marls) can therefore be considered as of late Eocene or early Oligocene age, with an absolute dating of about 35 million years BP (cf. Cavelier & Pomerol 1977).

Table 1 The age of the Bembridge Marls according to various authors, 1966-76

Oligocene: Lattorfian Edwards 1966 Oligocene: Lattorfian Curry 1966 Middle Oligocene: early Rupelian Rey 1967 Lower Oligocene . Castel 1968 Eocene : Ludian . Cavelier 1968 Oligocene: Tongrian Denizot 1968 Upper Eocene or Lower Oligocene. Curry, Gulinck & Pomerol 1969 Oligocene : Lattorfian Daley 1969 Oligocene: Tongrian Haskins 1971 Late Eocene-early Oligocene Keen 1972 Lower Oligocene . Martini 1972 Late Eocene or early Oligocene Bosma & Schmidt-Kittler 1972 Late Eocene or early Oligocene Bosma 1974 Early Oligocene Stinton 1975 Upper Eocene : late Priabonian Cavelier & Pomerol 1976 244 E. A. JARZEMBOWSKI Systematic descriptions The family and higher classification follows the papers of Gay, Riek and Common (all 1970) with the following departures: the Leuctrinae are regarded as a family following modern usage, a superfamily classification of the Neuroptera is not adopted because oflack of general agreement, and the classification of the Lepidoptera is also based on Karsholt & Nielsen (1976). Cockerell (l917b : 12) mentioned that he had described a species of Raphidia (order Megalop­ tera) from the Insect Bed; in fact (l917a : 373) he actually described a species of Riphidia (order Diptera : Tipulidae) which is a mis-spelling of Rhipidia Meigen 1818. A supplementary list of the material examined in the course of this study is deposited in the Palae­ ontology Library of the British Museum (Natural History). Except where otherwise indicated, the material is from the Insect Bed (Bembridge Marls) on the NW coast of the Isle of Wight, between West Cowes and the Newtown River. The figures prefixed '1.' or 'In.' are registration numbers of the Department of Palaeontology, British Museum (Natural History) (BM(NH)) and the material is currently housed in the Department of Entomology at the same instituti0- The major collections are abbreviated thus: S : E. J. A'Court Smith H : R. W. Hooley B : P. B. Brodie MIWG: Museum of Isle of Wight Geology Other collections are named in full. Wing veins are referred to by standard abbreviations; damaged or faint veins or margins a indicated by dashed lines, folds by dashed-and-dotted lines and extrapolations by dotted lines.

Order ISOPTERA Family MASTOTERMITIDAE

Genus MASTOTERMES Froggatt, 1896 TYPE SPECIES. Mastotermes darwiniensis Froggatt 1896 by monotypy; Recent, Australia.

Fig. 5 Mastotermes anglicus von Rosen. In.24571. (Scale line = 2 mm) PALAEOGENE INSECTS 245

Figs 6-7 AIastotermes anglicus von Rosen. Fig. 6, lectotype (hindwing), 1.15037. Fig. 7, paralecto­ type (forewing, 1.15033. (Scale line=2 mm)

Mastotermes anglicus von Rosen Figs 5-7, 15,20

1879 Phryganea F. Smith in Woodward: 344. 1913 Mastotermes anglicus von Rosen: 322; pI. 27, figs 5-8. 1965 = Mastotermes batheri von Rosen 1913; Emerson: 29. EMENDED DESCRIPTION. Forewing: elongate, some incomplete specimens suggesting that a greater elongation is possible than in the specimen figured by von Rosen (1913 : pI. 27, fig. 7). Sc short and single. Rl forking distad or basad of humeral suture, the outer branch terminating on the anterior margin at 0·2-0·4 of the forewing length (as measured from the suture). R2 and R3 elongate and separate at suture, R2 sometimes forked. Stem of Rs terminating near wing apex with a more or less marked anterior curve in its distal part. Rs with about four branches to the anterior margin, the outer ones usually subdivided. M and eu separate at suture, the former closer to Rs than euA with one to three major branches distally and perhaps several short branches towards base. In the left forewing of In.17143 and right forewing of 1.8659, M is briefly linked with Rs near base. Rs and euA are closer in some specimens than in von Rosen's figure with M nearly median. euA occupies about half of the wing area, dividing some ten times, the branches frequently forking once and occasionally several times. Humeral suture arcuate with maximum apical extension near M, and the basal part of the forewing is usually downcurved anterior to Rs. A brown pigmentation is often preserved on the scale, parts of M and on the more anterior veins. Forewings are preserved in 24 specimens and probably in two others. Hindwing: elongate, rounded apically; anterior and posterior margins nearly parallel, apex nearer the latter. Sc single, elongate, on average about five times as long as in the forewing; some faint branches to the anterior margin may be present. R1long terminating at 0·6 of the wing length from base, with none to four subparallel branches. R 2 + 3 has a common stem with Rs and 246 E. A. JARZEMBOWSKI

Sc+R Rs R, ~RS M

M-----::::::

Cu

11a 11b Figs 8-13 Kalotermes disruptus (Cockerell) comb. nov. Fig. 8, In.64534 (forewing). See also Fig. 17. Fig. 9, In.24619 (? hindwing). Fig. 10, In.24591 +In.24600 (hindwing). Fig. lla, 1.9845 (right forewing base). Fig. 11 b, 1.9845 (left forewing, anterior part of scale). Fig. 12, 1.9576 (hindwing base). Fig. 13, 1.8644 (holotype). See also Fig. 19. Fig. 14 Termitidae species A, In.24603. See also Fig. 35. Fig. 15 Mastotermes anglicus von Rosen, In.24610 (hindwing base). (Scale line= 2 mm. Figs 8-14 same scale, minor reticulation and membranous irregularities omitted). the appearance of being the first branch of the latter. Stem of Rs more or less oblique towards apex with two to four unforked major branches to the anterior margin. Stem of M nearer Rs than euA, united with Rs at base, and producing two to three major branches to the apical part of the posterior margin. Some very short branches are developed towards the base of M. CuA dividing ten to eleven times, the branches becoming progressively shorter towards base. CuP single with a broad terminal fork. lA parallel to anal fold with numerous short branches directed obliquely towards the latter. Anal lobe unknown, probably because it is folded under or broken away. The stem of M and the more anterior veins may show traces of brown pigmentation. As in the forewing, there is a well-developed reticulation. In addition to the lectotype, the hindwings are seen in six specimens and probably in two others. Body: about 15 mm long and all parts pigmented brown as in Recent Mastotermes. Head with a well-defined Y -shaped suture. Antennae about 5·0 mm long, segments transversely oval near base, circular near middle and elongate distally. Pronotum widest anteriorly with sides strongly PALAEOGENE INSECTS 247 downcurved (Fig. 20, p. 249). In.17357 has a longitudinal groove on the underside of the abdomen near the lateral margins as in Recent Mastotermes. In the same specimen, sternite 7 is not enlarged, 'ndicating that it is a male. Body remains are preserved in 17 specimens.

LECTOTYPE. 1.15037 (S), designated herein. Fig. 6.

OTHER MATERIAL. There are 18 paralectotypes and 42 other specimens (including one from St Helens), plus three possible specimens.

DIMENSIONS. Lectotype: wing length (apex-approximate position of 'suture') 25·8 mm, width 8·8mm. Other material. Forewing: length from humeral suture to apex (single specimen) 20·8 mm, maximum width 7·8-8·8 mm, length of Sc 1·0-2·8 mm, maximum length of scale 2·8-3·2 mm. Hindwing: length from wing base to apex 27·2-29·1 mm, maximum width 8,5-8,8 mm. Body: head, length 3·2-4·0 mm, maximum width 2·8-3·2 mm; pronotum, length 1·9-2·0 mm, maximum width 2,8-3,8 mm.

REMARKS. Emerson (1965 : 30) states that he examined the 'holotype' and 'paratypes' ofM. anglicus and that the species was described from three specimens. Examination of the fossil insect collection at the BM(NH) showed that three specimens of this species, labelled 'holotyp=' (1.15037) and 'paratype' (1.15033 and 1.8989), had been singled out and in addition there were a further 18 specimens each labelled 'paratype'. Von Rosen (1913 : 321) stated that he had examined 21 specimens of M. anglicus, and the former three correspond to the figures of his pI. 27; a foot­ note (1913 : 335) says that these are 'nach den Typen', but no holotype is designated in the paper. In a reprint from von Rosen in the library of the Department of Palaeontology, BM(NH), the registration numbers of the 21 specimens are hand written in the margin; 1.15037 is indicated as 'type' and the remainder as 'paratypes'. Von Rosen's material is therefore a syntypic series and US037 is here designated as lectotype. However, only 18 paralectotypes are recognized, as there are two pairs of part and counterpart; a further 42 specimens from the Insect Bed are also referred to this species. Eight specimens from the A'Court Smith collection, purchased in 1883 (1.15034-6, I.lS038-42) are labelled 'Phryganea', with or without a roman numeral in the same unidentified hand. Woodward (1879) recorded eight specimens of the trichopterous genus Phryganea in the A'Court Smith collection, on the basis of which a predator and prey association with Branchi­ podites vectensis was inferred by Tasch (1969). This misidentification was, however, realized not long afterwards, for on 1.8804 Brodie had written 'Phryganea' which he then crossed out and replaced by 'termite'; the specimen was purchased in 1898, a year after his death. The lectotype, a single hindwing, is adequately figured by von Rosen (1913 : pI. 27, fig. 8) except that Sc probably continues apically for twice the figured length but is intertwined with the fine anterior branches from R I . Both the wing and body remains ind